THE RAFFLES BULLETIN OF ZOOLOGY 2009
THE RAFFLES BULLETIN OF ZOOLOGY 2009 Supplement No. 20: 283–290
Date of Publication: 31 Jul.2009© National University of Singapore

ON THE IDENTITY OF PINNIXA BREVIPES H. MILNE EDWARDS, 1853, AND
A NEW SPECIES OF APHANODACTYLUS TESCH, 1918 (CRUSTACEA:
DECAPODA: BRACHYURA: PINNOTHEROIDEA) FROM THE PHILIPPINESPeter K. L. Ng
Tropical Marine Science Institute and Department of Biological Sciences,
National University of Singapore, Kent Ridge,Singapore 119260, Republic of Singapore
Email: dbsngkl@nus.edu.sg

Tohru Naruse
Transdisciplinary Research Organization for Subtropical and Island Studies,
University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903-0213, Japan
Email: naruse@lab.u-ryukyu.ac.jp

ABSTRACT. – A new species of Aphanodactylus is described from Panglao Island, the Philippines. This
new species can be distinguished from other species of Aphanodactlyus by the characters of the carapace,
the form of the ambulatory legs, and the structure of the male first gonopod. The photograph of the holotype
of poorly known Pinnixa brevipes H. Milne Edwards, 1853, is compared with species of Aphanodactylus
sensu stricto and the allied genus Gandoa Kammerer, 2006. Its distinctly punctate and rectangular carapacesuggest that A. brevipes belongs in Gandoa. A lectotype for Pinnixa brevipes is designated.
KEY WORDS. – Pinnotheridae, Aphanodactylus, new species, Philippines, taxonomy.

INTRODUCTION
Species of pinnotheroid crabs of the genus Aphanodactylus
Tesch, 1918, are known commensals of tube-dwelling
terebellid polychaetes of the genus Loimia. Aphanodactylus iscurrently represented by four species: A. sibogae Tesch, 1918
[type species; Lesser Sunda Islands], A. brevipes (H. Milne
Edwards, 1853) [Madagascar], A. edmondsoni Rathbun, 1932
[Hawaii], and A. loimiae Konishi & Noda, 1999 [Ryukyu
Islands] (Ng et al., 2008). The genus has traditionally beenplaced in the pinnotherid Asthenognathinae, but Ng et al.
(2008) recently transfered Asthenognathinae to Varunidae
and argued that Aphanodactylus and Gandoa Kammerer,
2006, a senior homonym of Voeltzkowia Lenz, 1905, willprobably need to be transferred to their own family at a later
date. A new genus and new species from Japan, closely allied
to these two genera has also been described by Naruse et al.
(2009). The establishment of this new family is now being
done by S. T. Ahyong and P. K. L. Ng, together with thedescription of a new genus from Guam.
The PANGLAO 2004 and PANGLAO 2005 Expeditions in
the central Philippines have yielded a bonanza of interesting

and rare species of decapod Crustacea, with dozens of new
genera and species reported thus far (see Bouchet et al.,2009; Ng et al., 2009; Richer de Forges et al., 2009). During
the PANGLAO 2004 expedition, one ovigerous female ofan unidentified species of Aphanodactylus was collectedfrom Panglao Island. In addition to this material, Johann
Hinterkircher and Arthur Anker kindly passed us a malespecimen of Aphanodactylus collected from a terebellid
worm at Panglao Island. In this paper, we report on a new
species of Aphanodactylus from Panglao Island.
The measurements provided, in millimetres, are carapace
width (CW) and length (CL) respectively. The abbreviations
P2–P5, G1 and G2 are used for pereopods 2–5, male firstand second gonopods, respectively. Specimens examined
are deposited in the Zoological Reference Collection (ZRC)of the Raffles Museum of Biodiversity Research, National
University of Singapore; Natural History Museum andInstitute, Chiba, Japan (CBM); Muséum national d’Histoire
naturelle (MNHN), Paris, France; Bernice P. Bishop
Museum (BPBM), Honolulu, Hawaii; and The Naturalis(ex Rijksmuseum van Naturrlijke Historie, RMNH), Leiden,
The Netherlands.

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Ng & Naruse: New species of Aphanodactylus from the Philippines
about two-thirds of ischium, concave distally; palp attachedto distolateral angle of merus, each segment not enlarged,
segments joined end to end; exopod slender, reachingproximal half of merus; flagellum long, reaching mesial
margin of merus.

TAXONOMY
Pinnotheridae De Haan, 1833
Aphanodactylus Tesch, 1918
Aphanodactylus panglao, new species(Figs. 1–4)
Aphanodactylus sibogae – Cases & Storch, 1981: 24, Fig. 9 (notAphanodactylus sibogae Tesch, 1918)
Material examined. – Holotype: ovigerous female (8.8 × 5.0 mm)(NMCR), Napaling, Panglao Island, Bohol Sea, Philippines, Panglao
Marine Biodiversity Project, PANGLAO 2004, stn. B8, 3 m deep,
9°37.1N 123°46.1E, by subtidal brushing of coral rock and debris, 7
Jun.2004. Paratype: 1 male (4.2 × 3.1 mm) (ZRC), Panglao Island,
Bohol Sea, the Philippines, associated with an unidentified terebellid
worm, coll. J. Hinterkircher, early 2000s.
Comparative material. – Aphanodactylus sibogae Tesch, 1918:
1 lectotype female (8.8 × 5.4 mm) (RMNH 2162), in tubes ofterebellid worm (Loimia), east of Dangar Besar, Sapeh Bay, northcoast of Sumbawa, Lesser Sunda Islands, Indonesia, up to 36 m
depth, coll. SIBOGA Expedition, 14–16 Feb.1900; 1 male (7.8 ×
5.3 mm) (ZRC), Sekotong, West Lombok, Indonesia, coll. D. L.
Rahayu, 16 May 2007. Aphanodactylus edmondsoni Rathbun, 1932:
1 holotype female (16.1 × 9.5 mm) (BPBM 3576), Oahu, Hawaii,coll. C. H. Edmondson, 27 Nov.1931; 1 male (11.6 × 8.0 mm)(BPBM 3577), Oahu, Waimanalo, Hawaii, coll. C. H. Edmondson,
27 Nov.1931; 1 male (11.5 × 8.2 mm), 1 female (16.9 × 10.2 mm)(ZRC 2000.0542), Oahu, Hawaii, coll. C. H. Edmondson, 1930s.
Aphanodactylus loimiae Konishi & Noda, 1999: 1 male (10.3 ×
7.4 mm), 1 ovigerous female (13.9 × 8.6 mm) (CBM-ZC 5341),
Kyan, Kuroshima Island, Yaeyama Islands, Ryukyus, Japan, 10 m
deep, coral reef, in tube of Loimia ingens, 10 Oct.1999, coll. K.
Nomura on SCUBA; 1 ovigerous female (15.3 × 9.1 mm) (CBM-ZC
5443), Ahra Beach, Kume-jima Island, Okinawa Islands, Ryukyus,
Japan, 10 m deep, coral reef, inhabited in tube of Loimia ingens,
15 Jun.1995, coll. K. Nomura on SCUBA.

Thoracic sternites 1/2 completely fused, 2/3 and 3/4
each demarcated by shallow ridge; sternites 4–8 laterally
demarcated by shallow sutures, each medially interrupted;
sutures between sternites 4/5 and 6/7 longer than others,
vulva on distal two-thirds of sternite 5, between mesial ends
of suture 5/6 and 6/7.
Chelipeds equal. Merus triangular in cross section, dorsal
margin rugose, dorsal margin and outer surface setose. Carpus
smooth, inner angle absent. Chela with smooth surfaces, palm
about 2.5 times longer than immovable finger; immovable
finger with weakly sinuous cutting edge, cutting edge convex
medially, lined with small teeth; movable finger with cuttingedge concave on distal half. Ambulatory legs (P2-P5)
relatively short, P3 longest, fourth leg shortest, upper surface
glabrous, setose on margins and lower surfaces of carpi andpropodi. Merus as long as or slightly longer than combined
length of respective propodus and dactylus, extensor marginunarmed, flexor margin with 0–2 small denticles (Table 1). All
propodi with 2 and 1 short, sharp claws on upper and lower
sides of distoflexor angle. Dactyli very short, claw-like.
Abdomen (Fig. 3h) with all somites and telson distinct, mobile;
somites wide, exopods of third to fifth pleopods protrudingfrom lateral margins of abdomen in ventral view.
Description of male paratype. – Carapace (Fig. 2)
subquadrate, broader than long, carapace width to length
ratio 1.35; dorsal surface smooth, regions poorly demarcated,

Description of female holotype (ovigerous). – Carapace (Fig.
1a) oval, distinctly broader than long, carapace width to length
ratio 1.76; dorsal surface smooth, regions poorly demarcated,pits absent from gastric region. Front (Fig. 1a) deflexed,
medially concave in dorsal view, gently convex in anteriorview. Orbital margins entire, slightly narrowed subdistallywith infraorbital and suborbital margins continuing laterally
beyond cornea to form relatively long gap; infraorbital margin
mesially ending as triangular inner orbital tooth; infraorbital
margin lining low suborbital crista, suborbital crista mesiallyreaching basal antennal articles. Anterolateral margin entire,
convex laterally, not cristate, continuous with posterolateral
margin. Epistome (Fig. 1b) very short, medially sunken.
Antennule folding transversely in fossa. Antennae withstout basal antennal article, not reaching distolateral angleof carapace; antenna enters orbit. Eyes (Fig. 1b) subconical,
eyestalk short, cornea distinct.
Third maxillipeds covering approximately three-quartersof buccal cavern when closed; ischium longer than merus,
extended proximo-laterally, with small protrusion on
distomesial angle; merus smaller than ischium, mid-length284

Fig. 1. Aphanodactylus panglao, new species. Holotype, ovigerous
female (8.8 × 5.0 mm) (NMCR): a, dorsal view; b, anterior view.

THE RAFFLES BULLETIN OF ZOOLOGY 2009
in female. Eyes (Fig. 3a) short, stout, with distinct cornea.
Third maxillipeds (Fig. 3b) covering almost whole buccal
cavern when closed; basically similar to female third
maxilliped except for more rounded distomesial angle.
Thoracic sternites 1/2 fused, sternites 2/3 demarcated by
distinct suture, 3/4 demarcated by low ridge, sternites 4–8
laterally demarcated by shallow, narrow sutures, sutures
medially interrupted, no medial longitudinal suture, press
button of abdominal locking mechanism present on distal
quarter of sternite 5; penis emerging from near anteriorborder of sternite 8 (Fig. 3e).

Fig. 2. Aphanodactylus panglao, new species. Paratype male
(4.2 × 3.1 mm) (ZRC). Arrows indicate low swellings on hepatic
regions.

with 2 pairs of small rounded swellings on hepatic regions.
Front (Fig. 3a) deflexed, produced triangularly in frontal
view, medially concave in dorsal view. Orbital margins (Fig.
3a) entire, oval, no gap between cornea and external orbital
angle, infraorbital margin mesially ending as triangular
inner orbital tooth, infraorbital margin lining low suborbitalcrista, suborbital crista mesially reaching basal antennal
articles. External orbital angle directed anteriorly, cristatefrom behind angle, anterolateral margin entire, continuous toposterolateral margin. Epistome (Fig. 3h) longer than that offemale, posterior margin straight. Antennule and antennae as
Table 1. Dentition on extensor and flexor margins of meri of P2–P5
of female holotype of Aphanodactylus panglao, new species.

P2 extensor margin
P2 flexor margin
P3 extensor margin
P3 flexor margin
P4 extensor margin
P4 flexor margin
P5 extensor margin
P5 flexor margin

Left

Right

0
1 large tooth0
2 large teeth
0
2 large teeth
0
0

0
0
0
2 large teeth
0
2 large teeth
00Chelipeds relatively strong, slightly unequal in size. Merus
triangular in cross section, ventro-outer margin granulated,dorsal margin rugose, dorsal margin and outer surface setose.
Carpus smooth, inner angle rounded. Chelae (Fig. 3c) andambulatory legs as in female, except for armature of extensor
and flexor margins of ambulatory meri (Table 2).
Abdomen (Fig. 3f) with all somites and telson distinct,
mobile, relatively narrow, first and second somites short,
third somite to telson forming straight lateral margins. G1
simple, proximal four-fifths straight, distal one-fifth bent
inwards towards median part of sternum.
Colouration. – Male dirty white; female pinkish-white.
Ecological note. – The holotype female was obtained fromcorals that were hand-collected by SCUBA divers. In thiscollecting method, divers brush coral rocks and debris,collecting the dislodged material in trays. Many of therocks are encrusted with terebellid worm tubes and thesewere probably destroyed during the brushing. The paratypemale was obtained from a terebellid worm collected by adiver (Fig. 4).
Etymology. – The species is named after the island fromwhich it was collected, Panglao. The name is used as a noun
in apposition.
Remarks. – Species of Aphanodactylus sensu stricto beara strong superficial resemblance to each other but can be
distinguished by a combination of characters. Aphanodactylus

Table 2. Dentition on extensor and flexor margins of meri of P2–P5 of male paratype of Aphanodactylus panglao, new species.
Left
P2 extensor margin
P2 flexor margin
P3 extensor margin
P3 flexor margin
P4 extensor margin
P4 flexor margin
P5 extensor margin*
P5 flexor margin*Right

7 large teeth with small denticles on proximal 1/2~3 large and small teeth on distal 2/3
With denticles, but very small
4 large teeth
Very small denticles
3 large teeth
0 or small denticles only
0 or small denticles only*

~6 large teeth with small denticles on proximal 3/5
2 large and some small teeth on distal 3/5
5 large and many small teeth on proximal 3/5
3 large teeth
Very small denticles
2 large with smaller ones
0 or small denticles only
0 or small denticles only

Many of male legs are detached, but on the basis of relative sizes, they are from both sides of P2 and right P3. Both P5 are missing, buton the basis of the photograph of the fresh specimen (Fig. 4), both margins appear to be unarmed or only with denticles.

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Ng & Naruse: New species of Aphanodactylus from the Philippines

Fig. 3. Aphanodactylus panglao, new species: a, cephalothorax, anterior view; b, left third maxilliped; c, right chela; d, left first ambulatoryleg; e, thoracic sternum; f, abdomen and telson; g, left G1, ventral view; h abdomen and telson. a–g, paratype male(4.2 × 3.1 mm) (ZRC);h, holotype, ovigerous female (8.8 × 5.0 mm) (NMCR). Scales bars: a, c–f, h = 1 mm; b, g = 0.5 mm.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009
panglao, new species, is distinct in the female possessing
relatively more slender ambulatory legs and the male having
a relatively narrower carapace (CW to CL ratio 1.35 versus
1.39–1.47 in the other species). In addition, the orbit of thefemale holotype of A. panglao is laterally long and slightly
narrowed sublaterally, and a relatively long gape is present
between external orbital angle and the tip of cornea (Fig. 1b).
In A. sibogae and A. loimiae, the orbit is gradually narrowed
laterally, and the gape between external orbital angle and thetip of cornea is proportionately shorter (Fig. 5b; Konishi &Noda, 1999: Fig. 2c). The extensor margins of the meri of
P2 to P5 of A. panglao are armed with small denticles orteeth, with that on the first ambulatory leg (P2) very strong,
consisting of six or seven prominent teeth (Fig. 3d). These
teeth are absent on A. edmondsoni and A. loimiae. They arealso likely to be absent or at best very small on type male A.
sibogae, with the detailed description and figures by Tesch(1918) showing an unarmed extensor margin of the merusof the third ambulatory leg. There are certainly no teeth or
distinct denticles on the extensor margins of the P2 to P5
meri of the present male of A. sibogae from Lombok. The
differences in the dentition observed in the ambulatory meral
flexor margins of the male and both margins in the case of
the female, do not appear to be significant. In addition, thehepatic region of the carapace of A. sibogae has two pairs ofelongated pits whereas in A. panglao, the pits are replacedby small swellings instead. Aphanodactylus panglao can also
be distinguished from A. loimiae and A. edmondsoni by its
distally bent male G1 (Fig. 3g) (versus gently incurved in

A. edmondsoni and A. loimiae) (Edmondson, 1962: Fig. 2d;
Ng et al., 2009).

Fig. 4. Aphanodactylus panglao, new species, paratype male (4.2 ×
3.1 mm) (ZRC), on unidentified host terebellid worm. Inset: freshlycollected male. (Photograph: J. Hinterkircher).

Fig. 5. Aphanodactylus sibogae Tesch, 1918, lectotype, ovigerous
female (8.8 × 5.4 mm) (RMNH 2162): a, dorsal view; b, ventral
view.

Cases & Storch (1981) recorded A. sibogae from Loimia
medusa in the Camotes Islands, Philippines. The number of
specimens and sexes examined were not listed, and the sizeand sex of the specimen figured was not specified; although
its appearance suggests it was a male. The figure of thismale, however, differs from males of other Aphanodactylus
species in having the anterolateral margins apparently morelaterally produced, relatively more convergent posterolateral
margins, and the fourth ambulatory leg being proportionately
stouter (Fig. 2; Cases & Storch, 1981: Fig. 9; Tesch, 1918:
Pl. 18 fig. 2, 2a; Ng et al., 2009: Fig. 1a). However, most of
the figures of decapod shrimps and crabs in Cases & Storch(1981) are rather schematic and not very accurate, even forseveral well known species. This suggests that their figureof “A. sibogae” may also be inaccurate and the differences
observed may be real. Our attempts to find the specimen(s)of “A. sibogae”, which are supposed to be in the biologicalcollections of the University of San Carlos, were unsuccessful.
The specimens are not in the university. Since Camotes isgeographically very close to Panglao Island (type localityof A. panglao), their record is provisionally also referred
to A. panglao.
Fransen et al. (1997) listed a syntype female of A. sibogae in
RMNH and this specimen was examined and photographed(Fig. 5). Tesch (1918: 285) stated that the size of the ovigerous
female is 11.3 × 6.0 mm, but the specimen on hand, however,measured 8.8 × 5.4 mm. It would appear that the original
measurements are in error. In any case, only two specimensof A. sibogae are known, one male, one ovigerous female,with the latter clearly a type (Fransen et al., 1997: 130).
The specimen agrees very well with Tesch’s (1918) original287

Ng & Naruse: New species of Aphanodactylus from the Philippines
description and figures. The only other type specimen,
the male, is still in the Zoological Museum, Universityof Amsterdam. The female specimen in RMNH is here
designated the lectotype of A. sibogae Tesch, 1918.

Gandoa Kammerer, 2006
Gandoa brevipes (H. Milne Edwards, 1853)(Figs. 6, 7)
Pinnixa brevipes H. Milne Edwards, 1853: 220; A. Milne-Edwards,
1873: 320; Tesch, 1918: 284 (footnote).
Aphanodactylus brevipes – Serène, 1964: 277, 278; Ng et al.,
2008: 247.
Material examined. – Lectotype female (ca. 8.5 × 5.1 mm, carapacesoft) (MNHN B-10616), Mayotte, coll. Cloué.

Remarks. – The taxonomy of this species is problematic.
In describing this species, all H. Milne Edwards wrote was“Carapace piquetée et sans crête transversale drolte maisprésentant vers le tiers postérieur un sillon courbe. Pattes
courtes et paraissant obtuses au bout, les dactylopoditesétant rudimentaires. Madagascar.” (H. Milne Edwards, 1853:
220). No figure was provided and he also did not indicate thenumber, sexes or sizes of the specimen(s) he had. AlphonseMilne-Edwards (1873), in naming the pinnotherid Pinnixa
fischeri (now in Tetrias) noted that in P. brevipes, “… sacarapace finement ponctuée et les pinces moins comprimées
et moins granuleuses.” (A. Milne-Edwards, 1873: 320). Tesch(1918: 284) also briefly discussed P. brevipes, noting thatits very short ambulatory dactylus was very similar to thatof Aphanodactylus and alluding to their close relationship.
Serène (1964) examined the type specimen of P. brevipes andcommented “Un examen du type femelle ovigère de 4 × 5
environ, confirme la supposition de Tesch (1918) au sujet deson appartenance au genre Aphanodactylus. En particulier les
mxp 3 sont sur P. brevipes comme décrits sur Aphanodactylus
sibogae : ischium trapezoide plus long que le merus, la suture

Fig. 6. Gandoa brevipes (H. Milne Edwards, 1853), lectotype female(ca. 8.5 × 5.1 mm, carapace soft) (MNHN B-10616): a, dorsal view;b, anterior view. (Photograph: J.-F. Dejouannet)

288

entre eux transverse, le palpe à 3 articles bout à bout, etc. … denombreux autres caractères concordent sur les deux espèces.
Cependant brevipes a les pattes ambulatoires beaucoup plus
longipes comparativement à celles de A. sibogae et c’est uneespèce differente qui justifie une nouvelle description avec
figure d’apres le type.” (Serène, 1964: 278). No photographor figure was provided. Serène (1964) clearly considered it
to be a species of Aphanodactylus, an action followed by
Ng et al. (2008).
Danièle Guinot was kind enough to check the type specimenof Pinnixa brevipes for us. She notes that the type is a driedovigerous female in poor condition, being soft and veryfragile. The label associated with it reads “Type. Mayotte,
Cloué” and appears to have been written by R. Serène. The
locality data, Mayotte, contradicts that stated by H. Milne
Edwards (1853), Madagascar, but this may not be significant.
The island of Mayotte is part of the Comoros, and like
Madagascar, used to be French territory. Mayotte lies justoff the northwest coast of Madagascar in the Mozambique
Channel. The identity of the supposed collector, Cloué, is
not known. As H. Milne Edwards (1853) did not select aholotype (or even indicate how many specimens he had), thefemale specimen in MNHN is here designated the lectotypeof Pinnixa brevipes.
The present photographs of the type of P. brevipes (Figs. 6,7) are the first of the species. As H. Milne Edwards (1853:
220) originally had described, the dorsal surface of thecarapace is clearly punctate, a feature not observed in theFig. 7. Gandoa brevipes (H. Milne Edwards, 1853), lectotype female(ca. 8.5 × 5.1 mm, carapace soft) (MNHN B-10616): a, ventral view;b, third maxillipeds. (Photograph: J.-F. Dejouannet)

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other three known Aphanodactylus species or the new species
here. In addition, Pinnixa brevipes has a characteristicallymore rectangular carapace (versus distinctly ovate in other
Aphanodactylus species; Figs. 1a, 5; Tesch, 1918: Pl. 18: Fig.
2; Edmondson, 1962: Fig. 3; Konishi & Noda, 1999: Figs. 1a,2a) and a laterally more elongate orbit (versus less elongatedin other Aphanodactylus species; Fig. 1b). These features
suggest that Pinnixa brevipes is closer to the monotypic
Gandoa Kammerer, 2006. Although Gandoa zanzibarensis(Lenz, 1905) [type locality: Zanzibar] is also a poorly
known species, with no additional record since the originaldescription, Lenz’s (1905: Fig. 9, 9a–c) drawings of the 8 ×
5 mm female clearly show the punctate dorsal surface of thecarapace and the laterally elongate orbit. Interestingly, both
species are from the east coast of Africa in the western Indian
Ocean, while Aphanodactylus is only known from the western
Pacific. The available evidence thus strongly indicates that
Pinnixa brevipes H. Milne Edwards, 1853, is not a species of
Aphanodactylus but should instead be transferred to Gandoa.
This action is taken here. Gandoa brevipes also appears to
be a different species from G. zanzibarensis, with the front
less well developed and the carapace proportionately muchbroader (cf. Figs. 6, 7; Lenz, 1905: Fig. 9). We also note thatthe presently observed differences between Aphanodactylus
and Gandoa are not substantial, and more specimens fromthe Indian Ocean may well result in the synonymy of thetwo genera. In view of the carapace differences and disjunctdistribution, we prefer to recognize Gandoa as a separate
genus for now.

thank our many colleagues who helped in the field collections,notably Marivene Manuel-Santos (NMCR) who was very
active in the intertidal collections. This study was supported
by research grant R-154-000-334-112.

ACKNOWLEDGEMENTS

Kammerer, C. F., 2006. Notes on some preoccupied names in
Arthropoda. Acta Zootaxonomica Sinica, 31(2): 269–271.

Thanks are due to Arthur Anker and Johann Hinterkircher forthe photographs and information of the male Aphanodactylus,
and entrusting the specimen to us for study.The male paratype
was collected during the Philippine-French-Singapore
Expedition, PANGLAO 2004, to the Bohol Sea, and we thankthe main organiser, Philippe Bouchet, and his Philippine
counterpart, Danilo Largo, for their kind help and support.
We gratefully acknowledge the strong financial support of
the TOTAL Foundation for this major effort, as well as the
support of the ASEAN Regional Centre for Biodiversity andConservation (ARCBC) in Manila, Philippines; Muséum
national d’Histoire Naturelle, Paris, France; and the National
University of Singapore. The strong support provided by
the University of San Carlos in Cebu, Philippines, notably
from Lawrence Liao in the pre-expedition period, is also
acknowledged. Thanks are also due to David Valdes(University of San Carlos) for his kind help in searchingfor the specimens of Cases & Storch (1981), and to J. C.
Mendoza for facilitating. Lu Eldredge kindly helped with
the Hawaiian material, and the first author acknowledges the
support from BPBM and him during an earlier visit. Thanks
are also due to Charles Fransen (RMNH) for his kind help.
Dwi Ristyo Rahayu kindly passed us a male specimen of A.
sibogae for this study. We also thank Danièle Guinot who
kindly checked the type of Aphanodactylus brevipes, while
Jean-François Dejouannet photographed it. Shane Ahyong
kindly helped with suggestions and some datasets. We also

Konishi, K. & H. Noda, 1999. A new species of commensal crab
genus Aphanodactylus (Crustacea: Brachyura: Pinnotheridae)
from the Yaeyama Islands, southern Japan. Publications of the
Seto Marine Biological Laboratory, 38(5/6): 223–229.

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